What makes animal gaits so audibly rhythmic? To answer this question, we recorded the footfall sound of 19 horses and quantified the rhythmic differences in the temporal structure of three natural gaits: walk, trot, and canter. Our analyses show that each gait displays a strikingly specific rhythmic pattern and that all gaits are organized according to small-integer ratios, those found when adjacent temporal intervals are related by a mathematically simple relationship of integer numbers. Walk and trot exhibit an isochronous structure (1:1)-similar to a ticking clock-while canter is characterized by three small-integer ratios (1:1, 1:2, 2:1). While walk and trot both show isochrony, trot has a slower tempo and is more precise and accurate, like a metronome. Our results quantitatively discriminate horse gaits based on rhythm, revealing striking commonalities with human music and some animal communicative signals. Gait and vocal rhythmicity share key features, and the former likely predates the latter; we suggest this supports gait-based hypotheses for the evolution of rhythm. Specifically, the perception of locomotor rhythmicity may have evolved in different species under pressure for predator recognition and mate selection; it may have been later exapted for rhythmic vocal communication.

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